REVIEWS REVIEWS REVIEWS Who needs environmental monitoring? Gary M Lovett1*, Douglas A Burns2, Charles T Driscoll3, Jennifer C Jenkins4, Myron J Mitchell5, Lindsey Rustad6, James B Shanley7, Gene E Likens1, and Richard Haeuber8 Environmental monitoring is often criticized as being unscientific, too expensive, and wasteful. While some monitoring studies do suffer from these problems, there are also many highly successful long-term monitoring programs that have provided important scientific advances and crucial information for environmental policy. Here, we discuss the characteristics of effective monitoring programs, and contend that monitoring should be considered a fundamental component of environmental science and policy. We urge scientists who develop monitoring programs to plan in advance to ensure high data quality, accessibility, and cost-effectiveness, and we urge government agencies and other funding institutions to make greater commitments to increasing the amount and long-term stability of funding for environmental monitoring programs. Front Ecol Environ 2007; 5(5): 253–260 W e use monitoring data routinely in our daily lives; we monitor the stock market, the weather, our blood pressure, and baseball statistics. But, does monitoring have a place in environmental science? Common criticisms of environmental monitoring are (1) that it is not really science, but merely a fishing expedition that diverts funds from “real” science, (2) that most monitoring data are never used, and (3) that we can’t possibly know today what critical questions will need to be answered in the future. Some people feel that monitoring has no place in rigorous environmental science, and mindless monitoring gives the discipline a bad reputation. Who needs it? In a nutshell: • Environmental monitoring is often criticized as being unscientific, expensive, and wasteful • We argue that monitoring is a crucial part of environmental science, costs very little relative to the value of the resources it protects and the policy it informs, and has added value in that basic environmental monitoring data can be used for multiple purposes • Effective monitoring programs address clear questions, use consistent and accepted methods to produce high-quality data, include provisions for management and accessibility of samples and data, and integrate monitoring into research programs that foster continual examination and use of the data • Government agencies should commit to long-term support for valuable monitoring programs, and funders of basic ecological and environmental research should recognize that monitoring is a fundamental part of environmental science 1 Institute of Ecosystem Studies, Millbrook, NY *(lovettg@ecostudies.org); 2US Geological Survey, Troy, NY; 3Syracuse University, Syracuse, NY; 4University of Vermont, Burlington, VT; 5College of Environmental Science and Forestry, State University of New York, Syracuse, NY; 6US Department of Agriculture Forest Service, Cumberland, ME; 7US Geological Survey, Montpelier, VT; 8US Environmental Protection Agency, Washington, DC © The Ecological Society of America In this paper, we evaluate these common perceptions of environmental monitoring and discuss the characteristics of successful and unsuccessful monitoring programs. We define environmental monitoring as a time series of measurements of physical, chemical, and/or biological variables, designed to answer questions about environmental change. These measurements may be taken at one or multiple locations. The meaning of “long term” depends on the time scale of the ecological process of interest, but in this paper we focus on datasets that span decades. Our discussion is particularly relevant, given the budgetary constraints on current monitoring and the ongoing debate regarding the opportunities, limitations, and costs associated with the establishment of national environmental observatories in the US, for example the National Ecological Observatory Network (NAS 2004), the Hydrologic Observatories, and the Cooperative Largescale Engineering Analysis Network for Environmental Research (NRC 2006). Is monitoring science? Good science involves more than just devising clever experiments to test a specific hypothesis. Carpenter (1998) suggests that ecosystem science is like a table supported by the four legs of theory, experimentation, crosssite comparisons, and long-term studies. Many ecosystems require long-term study because they change slowly, and sustained monitoring of key variables provides the principal record of change. Long-term monitoring data also provide context for short-term experiments and observations. For example, suppose an elegant irrigation experiment is conducted to determine the effect of water availability on plant production, and no response is observed. If long-term measurements of precipitation indicate that the experiment occurred during the wettest year on record, the monitoring data provide important information for interpreting these unexpected experiwww.frontiersinecology.org 253 GM Lovett et al. Environmental monitoring mented, they also add little to scientific understanding. The keys to good science are similar for all forms of scientific inquiry, including monitoring: good questions, appropriate research designs, high-quality data, and careful interpretation of the results. 254 Courtesy of EPA Clean Air Markets Division Is monitoring cost-effective? Monitoring not only provides the basis for formulation of science-based environmental policy, but continued monitoring also permits evaluation of whether the policy has had its intended effect and has been cost-effective. Solutions to many environmental problems are expensive and technically challenging, but the cost of monitoring programs is generally much less than either the cost of policy implementation or the monetary benefits associated with environmental improvement. The costeffectiveness of monitoring in implementing environmental policy is illustrated below by three examples of major environmental issues and their Figure 1. The National Atmospheric Deposition Program (NADP) and related monitoring efforts. Clean Air Status and Trends Network (CASTNET) site at Prince Edward, VA. These two atmospheric monitoring networks are crucial for Clean Water Act quantifying atmospheric deposition and tracking responses to air-pollution legislation. The two tall towers and the instrument shed at the left of the The Clean Water Act (CWA) of 1972, along with photo hold the instrumentation for the air chemistry and dry deposition subsequent amendments, has established the modmeasurements made by CASTNET, and the automatic precipitation ern framework for regulating water pollution in the collector at the right of the photo is used by the NADP for measuring rivers, lakes, wetlands, and coastal waters of the precipitation chemistry. US. The CWA regulates discharge from municipal wastewater treatment plants and industrial facilimental results. Monitoring data are also necessary for ties, non-point source pollution from agricultural actividetermining whether an event is unusual or extreme and ties, wetlands protection, and many other activities. The are useful for the development of appropriate experimen- cost of water-quality monitoring under the CWA is diffital designs. For example, long-term monitoring of atmos- cult to estimate because of the diffuse responsibilities of pheric ozone provides the critical information on average federal, state, and local agencies under various sections of and extreme ozone concentrations that is needed to this complex law. Accordingly, estimates of the total costs design an appropriate experiment examining the effects of complying with the CWA vary widely, from $14.1 bilof ozone on plants. Monitoring data also contribute sig- lion per year (in 1997 dollars; Van Houtven et al. 2000) nificantly to the parameterization and testing of environ- to as high as $93.1 billion per year (in 2001 dollars; mental models, such as the general circulation models Johnson 2004). Judging the effectiveness of these expen(GCMs) that predict future climate change. Clearly, ditures in improving water quality has been contentious (Smith et al. 1987; Knopman and Smith 1993), due in monitoring is a crucial part of environmental science. Proposals that include monitoring generally do not fare part to concerns about the quality, statistical representawell in competition for basic science funding from com- tiveness, and comparability across states of long-term petitive grant agencies, such as the US National Science water quality measurements, which are largely the Foundation. Reviewers often object on the grounds that responsibility of individual states (Adler et al. 1993; GAO monitoring does not address a hypothesis, or that the 2000; Hirsch et al. 2006). A recent estimate of governhypothesis is trivial. However, careful, structured obser- ment expenses for regulation and monitoring under the vation is as fundamental to science as is hypothesis test- CWA puts the cost at $982 million in 2001, of which ing. Good, long-term monitoring records are rare and much less than half is likely to be attributable to waterpollution monitoring (Johnson 2004). If as much as oneextremely valuable. Haphazard monitoring, of course, adds little to scien- third (probably an overestimate) of these expenses are for tific knowledge. However, this also applies to other forms actual water-quality monitoring, then this value repreof scientific endeavor; when theory, experiments, or sents only 0.4–2.1% of the estimated cost of complying cross-site comparisons are poorly designed or imple- with the CWA. www.frontiersinecology.org © The Ecological Society of America GM Lovett et al. Environmental monitoring Acid deposition 255 Acid deposition has been an important scientific and policy issue in North America since the 1970s. Major policy action was taken in the 1990 amendments to the Clean Air Act, in which Title IV is focused on protecting ecosystems from acidifying deposition of sulfur and nitrogen. Title IV has resulted in major reductions in emissions of sulfur dioxide (SO2) and nitrogen oxides (NOx) by electric power generation plants. By 2005, Title IV emission reduction programs had diminished annual SO2 emissions by more than seven million tons from 1980 levels, and had reduced annual NOx emissions by over three million tons from 1990 levels. The most recent analysis of the benefits and costs of compliance with Title IV estimated the program’s benefits at Figure 2. US Department of Agriculture Forest Service technicians measure $122 billion annually in 2010, when the decaying wood in the Delaware River Basin as part of the Forest Inventory and program will be fully implemented, while Analysis (FIA) program. The FIA program monitors forest health and projecting the annual cost to industry of productivity throughout the US (www.fia.fs.fed.us/). compliance at approximately $3 billion (in 2000 dollars; Chestnut and Mills 2005). Changes in acid and, in addition, may be of future economic importance if deposition resulting from Title IV are primarily moni- the US adopts a carbon-trading program that includes tored by two networks (Figure 1): the National credits for carbon sequestration. The potential for US Atmospheric Deposition Program (NADP), which mea- forests to sequester atmospheric CO2 has been estimated sures precipitation chemistry in the US at a cost of from the data collected by the US Department of approximately $6 million annually (V Bowersox pers Agriculture Forest Service's Forest Inventory and comm), and by the Clean Air Status and Trends Network Analysis (FIA) program (Birdsey 1992; Birdsey and Lewis (CASTNET), which measures ozone and dry deposition 2003; Figure 2). The primary goal of the FIA program is at a cost of approximately $3.9 million annually. In addi- to monitor the extent, condition, productivity, uses, tion to these air-quality monitoring programs, the US impacts of management, and health of forest ecosystems Environmental Protection Agency (EPA) has two sur- in the US, but the FIA data are used by many researchers face-water monitoring programs to assess the response of for a wide variety of purposes, including estimation of foracid-sensitive watersheds to changes in atmospheric est carbon sequestration. Using FIA data and other infordeposition (Stoddard et al. 2003); they are the mation, Goodale et al. (2002) estimated that the annual Temporally Integrated Monitoring of Ecosystems (TIME) accumulation of carbon in forest ecosystems in the US and the Long-Term Monitoring (LTM) program, which from 1990–1991 was 0.28 Pg, or 280 million tons. The differ in intensity of sampling and selection of sampled monetary value of this ecosystem service is difficult to lakes and streams. The TIME/LTM program currently quantify. However, recent analyses of energy policy focuses on the areas most impacted by acid deposition: options suggest that if a carbon-trading program were lakes in the Adirondack Mountain region of New York implemented in the US, carbon credits could be priced at and New England and streams in the Northern $5 to $25 per ton (National Commission on Energy Appalachian Plateau and Blue Ridge region of Virginia Policy 2004, 2005), yielding a value for forest carbon and West Virginia. The scope and budget of this program sequestration of between $1.4 billion and $7 billion per has been markedly reduced, from $2 million annually in year. The federal appropriation for FIA in 2003 was $59.7 the early 1990s to substantially less than $1 million today. million, supplemented by $10.2 million from other partTaken together, the total cost of these critical atmos- ners (USDA Forest Service 2004). Thus, depending on pheric deposition and surface-water monitoring programs the price chosen for carbon credits, the cost of implerepresents less than 0.4% of the implementation costs of menting the FIA program ranges from < 1% to 5% of the Title IV and less than 0.01% of the estimated benefits. potential value of annual carbon sequestration in forest ecosystems in the US. This analysis is for carbon sequestration alone, and does not consider the many other Carbon sequestration important uses of FIA data, such as evaluation of forest The potential for vegetation in the US to sequester car- health and timber stocks. bon is important for the analysis of global carbon budgets The above examples show clearly that while large-scale © The Ecological Society of America www.frontiersinecology.org Environmental monitoring 256 390 380 CO2 (ppmv) 370 360 350 GM Lovett et al. tion work, the small fraction spent on monitoring actually provides an opportunity to learn which restoration methods work and which do not. Are monitoring data used effectively? 340 We all know of monitoring data that have accumulated in file cabinets (or on hard drives) and gathered dust. These 320 data may be either inaccessible to all but a few, too poor in 310 quality or too poorly documented to be useful, or they may 300 have been collected solely to fulfill a legal requirement, Jan 58 Jan 62 Jan 66 Jan 70 Jan 74 Jan 78 Jan 82 Jan 86 Jan 90 Jan 94 Jan 98 Jan02 with no real motivation for thorough analysis and interpreFigure 3. Mean monthly atmospheric concentrations of CO2 tation. At the other end of the spectrum are datasets from measured at Mauna Loa, HI, by CD Keeling and colleagues both individual investigators and large institutional profrom 1958 to 2004. Plotted from data available at the Carbon grams that have enormous value to environmental science Dioxide Information Analysis Center (www.cdiac.ornl.gov). and policy. Consider, for example, Charles David Keeling’s record of atmospheric CO2 concentrations at Mauna Loa, monitoring programs often require considerable expendi- Hawaii. Although Tyndall and Arrhenius proposed in the tures, in many cases these costs are miniscule compared late 19th century that changes in atmospheric CO2 conto the value of the resources monitored and the policies centrations could affect the heat budget and surface temthat affect these resources. Furthermore, the absence of perature of Earth, the concept of global climate change monitoring can greatly hinder evaluation of the effective- and its linkage with anthropogenic emissions of CO2 did ness of environmental policies and programs. One exam- not emerge until Keeling began long-term measurements ple is the lack of a uniform and integrated surface-water of atmospheric CO2 at Mauna Loa in the 1950s (Figure 3). monitoring program to assess the Clean Water Act, as Now, global climate change is arguably the foremost issue discussed above. Another is the inability to assess the in environmental science and policy. Consider also measurement of the chemistry of bulk preeffectiveness of the $14–15 billion spent on river restoration projects in the US since 1990, because these projects cipitation and streamwater at the Hubbard Brook rarely include river monitoring before and after the Experimental Forest in New Hampshire (Figures 4 and 5). restoration effort (Bernhardt et al. 2005). While it may Begun in 1963, this is the longest-running precipitation seem that monitoring costs take funds away from restora- and streamwater chemistry record in the US and has figured prominently in the development of the watershed–ecosystem concept, analysis of watershed bio160 geochemical cycles, and evaluation of forest Streamwater 140 harvesting policy (Bormann and Likens 1967; 120 Likens et al. 1978; Likens and Bormann 1995). During the scientific debate that preceded the Clean 100 Air Act Amendments of 1990, the sulfate trends 80 Precipitation (Figure 4) at Hubbard Brook were very influential in convincing scientists and policy makers that 60 decreasing sulfur emissions would produce substan40 tial decreases in sulfur deposition and streamwater sulfate concentrations in the northeastern US 20 (NRC 1983). 0 A good example of a valuable, broad-scale moni1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 toring program is the NADP precipitation chemWater year istry network discussed above. The NADP data are Figure 4. Long-term record of annual volume-weighted mean sulfate widely used because of their high quality and free concentrations in bulk precipitation and streamwater in Watershed 6 of availability on the Internet. These data have the Hubbard Brook Experimental Forest, NH (updated from Likens et proven effective in evaluating emissions policies al. 2002). This data record was initiated as part of a basic study of forest for sulfur and nitrogen oxides. For example, they ecosystem function, but has also proven valuable in assessments of were used to measure the decline in atmospheric ecosystem response to policies controlling emissions of air pollutants. nitrate and sulfate deposition following the impleMonitoring at the Hubbard Brook Experimental Forest relies on an mentation of the 1990 Clean Air Act effective collaboration between academic scientists and the US Amendments (Lynch et al. 2000; Butler et al. 2001, Department of Agriculture Forest Service, which monitors hydrological 2005). The NADP program also provides a powerand meteorological variables at this site. ful example of the value of the Internet in providSulfate concentration (µeq l-1) 330 www.frontiersinecology.org © The Ecological Society of America GM Lovett et al. Environmental monitoring ing broad-scale access to data. Prior to 1995, when the NADP data were first made available online, fewer than 500 people per year received data on tape or diskette (V Bowersox pers comm); in contrast, more than 18 000 downloads of NADP data from the Internet have been recorded annually in recent years (NADP 2003). Another crucial, large-scale monitoring program is the USGS national network of stream gauges, which provides fundamental data nation-wide for flood prediction and estimation of water supply. The data are used for other scientific purposes as well, including analysis of the impacts of climate change (Lins and Slack 1999; McCabe and Wolock 2002; Hodgkins et al. 2003) and testing the accuracy of models that predict future climate change Figure 5. The V-notch weir at Watershed 6 of the Hubbard Brook (Milly et al. 2005). The examples above are well known, but Experimental Forest, NH. The long-term stream chemistry monitoring data even relatively obscure datasets can be valu- shown in Figure 4 are from this stream. able if they are accessible and of high quality, and if the record is of sufficient length. Climate itoring data with enduring value. Good foresight involves researchers have unearthed many long-term records that looking ahead to anticipate potentially important enviprovide information on climate change. Examples from ronmental problems of the future and selecting key meathe US include the record of dates of spring-time wild- surements that are likely to be sensitive to change. flower blooming and bird arrivals that Aldo Leopold Understanding the system depends on choosing the most began at his Wisconsin farm in 1936 (Bradley et al. 1999), relevant measurements among a host of possibilities in the changing bloom dates of lilac planted at hundreds of complex systems. Some variables are important because sites in the mid-1960s (Schwartz and Reiter 2000; Wolfe they represent critical processes (eg net primary producet al. 2005; Figure 6), and the long-term records of lake tion, nutrient budgets); others are useful because they are ice-out dates (Likens 2000; Magnuson et al. 2000; known controllers of ecosystem function (eg precipitaHodgkins et al. 2002) and ice thickness of rivers tion amount, phosphorus concentration in lakes); still (Huntington et al. 2003). others are important because people care about them (eg The criticism that monitoring data are collected and bird species diversity). Monitoring such key variables never used is unfortunately true in some cases, but other improves the probability that a dataset will be useful for monitoring datasets have proven extremely important. the study of future environmental issues. Below, we offer some guidelines for maximizing the use and value of monitoring programs. 257 Can today’s monitoring programs Good monitoring programs are designed around good questions, but will the questions asked today still be important decades into the future? Unfortunately, no one knows for certain which environmental issues will emerge and which will recede as time passes. No doubt, some scientist of the next century, desperately in need of historical data to address an important problem, will look back and wonder how environmental scientists could have failed to take some crucial measurement, whatever it might be. However, a combina- Figure 6. The lilac variety Syringa chinensis, “red rothomagensis”, in full tion of good foresight and understanding of bloom. Flowering dates of clones of this plant are used for monitoring the system being monitored can produce mon- phenological trends associated with climate change throughout the US. © The Ecological Society of America www.frontiersinecology.org Courtesy of M Schwartz answer tomorrow’s questions? GM Lovett et al. Environmental monitoring 258 Panel 1. The seven habits of highly effective monitoring programs* (1) Design the program around clear and compelling scientific questions. Questions are crucial because they determine the variables measured, spatial extent of sampling, intensity and duration of the measurements, and, ultimately, the usefulness of the data. (2) Include review, feedback, and adaptation in the design. The guiding questions may change over time, and the measurements should be designed to accommodate such changes.The program leaders should continually ask,“Are our questions still relevant and are the data still providing an answer?” The program should have the capacity to adapt to changing questions and incorporate changing technology without losing the continuity of its core measurements. (3) Choose measurements carefully and with the future in mind. Not every variable can be monitored, and the core measurements selected should be important as either basic measures of system function, indicators of change, or variables of particular human interest. If the question involves monitoring change in a statistical population, measurements should be carefully chosen to provide a statistically representative sample of that population. Measurements should be as inexpensive as possible because the cost of the program may determine its long-term sustainability. (4) Maintain quality and consistency of the data. The best way to ensure that data will not be used is to compromise quality or to change measurement methods or collection sites repeatedly.The confidence of future users of the data will depend entirely on the quality assurance program implemented at the outset. Sample collections and measurements should be rigorous, repeatable, well documented, and employ accepted methods. Methods should be changed only with great caution, and any changes should be recorded and accompanied by an extended period in which both the new and the old methods are used in parallel, to establish comparability. (5) Plan for long-term data accessibility and sample archiving. Metadata should provide all the relevant details of collection, analysis, and data reduction. Raw data should be stored in an accessible form to allow new summaries or analyses if necessary. Raw data, metadata, and descriptions of procedures should be stored in multiple locations. Data collected with public funding should be made available promptly to the public. Policies of confidentiality, data ownership, and data hold-back times should be established at the outset. Archiving of soils, sediments, plant and animal material, and water and air samples provides an invaluable opportunity for re-analysis of these samples in the future. (6) Continually examine, interpret, and present the monitoring data. The best way to catch errors or notice trends is for scientists and other concerned individuals to use the data rigorously and often. Adequate resources should be committed to managing data and evaluating, interpreting, and publishing results. These are crucial components of successful monitoring programs, but planning for them often receives low priority compared to actual data collection. (7) Include monitoring within an integrated research program. An integrated program may include modeling, experimentation, and cross-site comparisons. This multi-faceted approach is the best way to ensure that the data are useful and, indeed, are used. *With apologies to Covey (1989) Observation of trends or spatial patterns in monitoring data often allows us to discover emerging environmental problems and to advance environmental science in new directions. Examples include Keeling’s measurements of increasing atmospheric CO2 (Keeling et al. 1976; Figure 3) and the discovery of the “ozone hole” above Antarctica by a team of British scientists (Farman et al. 1985). Each of these discoveries led to new research and policy efforts that have reverberated around the globe. Finally, scientists of the future will look back with profound gratitude if we archive samples and not just data. Chemical measurement techniques are sure to improve, and new problems will require analyses that have not yet been developed. When samples of soil, plant material, animal specimens, water, and other environmental media are archived, the scientists of the next century will have a treasure trove of stored information with which to answer the burning questions of the day. What makes a monitoring program successful? In this paper, we have highlighted examples of environmental monitoring that have proven useful in furthering science and policy. Yet, as we point out above, some monitoring datasets go largely unused. How can we ensure that monitoring programs, which consume our time and www.frontiersinecology.org financial resources, will be useful in the future? There is no single best model for the structure of a monitoring study. Effective monitoring has been done by individuals and small groups of scientists and by large agencies, and has occurred at individual sites, across large regions, and even over entire continents. Like other forms of long-term research, the most important requirement is a personal or institutional commitment to sustain the program (Strayer et al. 1986). When the goal is to maintain a consistent measurement regimen for decades, there will undoubtedly be times when funding will be lean and much effort will be required to continue the endeavor. A successful program must be designed to survive lean times by maintaining a solid funding base, a core set of inexpensive measurements, and a group of individuals dedicated to collecting, interpreting, and using the data. Beyond this personal or institutional commitment, successful monitoring programs have several important characteristics of design and implementation, which we summarize in Panel 1. Conclusions and recommendations Who needs environmental monitoring? We all do. Scientists need monitoring as part of integrated environmental research programs. Policy makers need monitor© The Ecological Society of America GM Lovett et al. ing to design, implement, and evaluate effective environmental policies. The public needs monitoring to track our nation’s natural resources. Monitoring is not second-rate science. Rather, it is an essential component of environmental science and deserves the careful attention of scientists and greater support from government agencies and other funding sources. If you are a scientist managing a monitoring program, your responsibility is to ensure that the data are of high quality, that data and methods are broadly accessible, and that the program is as cost-effective as possible. The “seven habits” listed in Panel 1 provide good operating principles for these programs. Greater commitment is needed to examine, interpret, and apply data that are being collected and to cooperate with other scientists in forming monitoring networks that are designed to promote comparability of data across sites and across scales of space and time. If you are a policy maker, resource manager, or a program manager in a government agency or funding institution, your responsibility is to make the commitment to maintain and expand long-term monitoring programs. Fickleness of funding has led to the demise of many good monitoring programs, and Herculean efforts are often required by scientists to continue collecting data in the face of reduction or loss of funds. Agencies such as the US Geological Survey, the US Department of Agriculture Forest Service, the National Oceanic and Atmospheric Administration, and the US Environmental Protection Agency should evaluate their monitoring programs in the light of national needs and take steps to ensure that those programs are not sacrificed in tight budget years. The National Science Foundation and other agencies with competitive grant programs need to recognize that longterm monitoring is a critical part of the infrastructure of environmental science and that many valuable monitoring programs are funded through investigator-initiated research proposals. Maintenance of long-term monitoring datasets should be a highly valued feature in the review of proposals. At a national scale, better integration of monitoring efforts is needed to make more efficient use of the funds now being spent on monitoring by many federal and state agencies, universities, and private companies (CENR 1997), and more funding is required to monitor critical environmental indices that are not currently being measured (Heinz Center 2006). A renewed commitment to funding environmental monitoring, and a clear focus on doing it well, will provide an invaluable legacy for future scientists and citizens. Acknowledgements This paper is a project of the Northeastern Ecosystem Research Cooperative. We thank R Skeffington for inspiration and D Sleeper for advice and encouragement on this paper. We acknowledge the support of the US National Science Foundation through its “Research © The Ecological Society of America Environmental monitoring Coordination Networks” program (grant DEB-0342198) and through grants DEB 0129138 and DEB 0423259. 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